植物科学最前線 3: 84 ( 2012)= ūǜȀǻŹŨŀōŀō̯ɁōɣɲɪɃȹɪōśƶ    ǡŹŨů̧ť 
An introduction to the taxonomy and morphology of Marchantia polymorpha
Key words: bryophytes; Marchantia polymorpha; morphology; taxonomy
Masaki Shimamura
Department of Biology, Graduate School of Science, Hiroshima University
Kagamiyama 1-3-1, Higashi-Hiroshima 739-8526, Japan
A  r~kh@͚üĀɀY§ƒ=¤ɇ§=èŰ+͚²̫?ÃɂĸĞ́9ǒOǍ̌=LSVUjhǱȹ?͜69UjhǱȹ=ȿ·ɒ<è͌ō!̘Ʌ)VU¸ò S͚Ƿ<ǚˀ˥ɹ=ɐĴ+8
T͚Ǳȹō?Ēáɒ<ƸɪǑ9@͚t^͌?·ːÇ:+8HIÇľ<#ďTƣXV8UwPɼī9Ē# SǱȹō?ƸʦɅ:+8Ź#ɅSV2 Leopold Kny =RU̎?Ǳȹ
ĩˬ (18741911)?§9͚ˤʉ<r~kh?ĩȸ!çȸ)V2':O (Kny1890)͚r~kh!t^͌?·ːÇ:+8ƣXV8"2ɁɆ9@< :ƪƳ)V8U (Stotler 1994)19 ¤ʃ=
8@͚O5:Oɣɲ?̏Z92Ǳȹ?͜69T͚{Ǳȹ?Ùɨ:OĠGUŋĭ9U
'VK9?̨ɣɲ?ȂĔ9ˇɰ+2ʱŀ<ɠ˙@͚iƘĳPèň̙½ō?ƠȒYɅ2ɣɲY̏NU¡9Oŀ"<˲Ʉ:<U:ʢSVU(ȏâɢũ 2012)ǚɯ9@͚r~kh=̬Uɣɲ?§9OǒOĒȂĔYO6è͌ō:Ɔƚō͚0?Ğ́è̡=̬-Uɠ˙YK:N͚³Ƌ
?ɣɲ?ŢǕYɧ+2B

=.29.7

 jhǱȹ@ʔɻǝYO2<̝ÐÄ!͚¨˗<Ƀȕ¤·9UȲ9ȺƑ7%SV͚t^͌
(Marchantiophyta)͚q͌ (Bryophyta)͚xkh͌ (Anthocerotophyta)?͞ʟ=è͌)V8U'VSʟ?̝ÐÄ@̨̏û?ȂĔ90V1V=ĿǷûY̐&8U2N͚̝ÐÄ?Ɔƚ9ʟ
Yè%UƆ˳YȺƑ7%U?@̺+!͚ʪňÄ?Ɔƚ!ȍŒɒ=̖58Uq͌:xkh͌?ʪňÄ!ȋŉYÔ8U?=Ś+͚t^͌@O58<K2͚t^͌?ʪňÄ@͚̝ÐÄɆǟ?Íˮģő=ŐÝ=˘XV2Ȼƚ9Ɯ̨+͚ȟƺè˓!ʋ¬+2Ƌ=ʪňÄ?ǧ!¿̨+͚@,N8ľ̛=̾ç-Uq͌?ʪňÄ@͚ͅɷ̛=̝ÐÄɆǟ?ʌʜ?̛͘ˌŴ͙
Y˒52Ȼƚ9Ɯ̨+͚ǧ!¿C8ʪňÄ!ľ̛=̾ç+2Ƌ=ʪňĥ!ɏ̕+͚ȟƺè˓!
'Uxkh͌?ʪňÄ@͚Ĳ̛=è˓ʌʜYO4͚¡ǂ=ę%8ǁ2<ʪňÄʌʜYʒʓɒ=ƆƜ+ʓ%Ut^͌:q͌9@ʪňÄâ̛?ʌʜèû!ė˩ɒ='T͚ʪňƆƜ͘ȟƺ

M. Shimamura-1
BSJ- Revi ew 3: 84 ( 2012)

植物科学最前線 3: 85 ( 2012)è˓͙OʪňÄâ̛9ėǌ=Ő¬-U!͚xkh͌9@͚ʪňÄ?ʌʜèû@Ĳ̛ S¡ǂ=
ę%8̊ǿɒ='T͚ʪňOǁ2<O?!̊ǿƆƜ)VU t^͚͌q͚͌xkh͌?ʂʐ̬Ë=68@Ƿ<˭˪!U!͚ǒ̃?èňʂʐōɒ˞ǣ?Ŀ#@͚r~kh!ěKVUt^͌!ǒOëǖ=èŦ+2':YƵƩ+8U (Qiu et al.
2006, 2007)ǒǁ?è͌Äʂ (Crandall-Stotler et al. 2009) 9@͚t^͌@͚ŀ"#jvkhʕ
(Haplomitriopsida)͚r~khʕ (Marchantiopsida)͚xkhʕ (Jungermanniopsida) ?͞ʟ=ŀí)V8Ujvkhʕ͘ʄ20ɮ͙@͚ǒOëǖ=èŦ+2:ʢSV8U͚ɗɶU@ùú-Uʿ˃Ä?ľƆYO6è͌ʟ9U̍ĆģP̍ʀģYÍˮ-U2N?Ⱥí<˃ɒģő
YO2<͚¹ǯYO2<<;͚jhǱȹ:+8OÇľɒ<ȺƑ!T͚t^͌?§9OĈŇ
ɒ<ƆƚYɈN8U:ʢSV8Uxkhʕ@t^͌?ŀĀ?ɮYěMè͌ʟ9͚0?ƆƚOĿǷ9T͚ɗɶ-Uʿ˃Ɩ͚ɃʦĲȹ=ȑ58ùú-Uʿ˃Ɩ͚ʿ:˃?èû!£Ǉ
ɞ9â̛ʌʜ?èûOĂʆ<Ăʆ˃ȻƖ (simple thalloid):Ƿ9Ur~khYěMr~khʕ͘ʄ400ɮ͙@͚ƞŷ<˃ȻÄ?ľƆYO6g9Ur~khʕ9@͚ľɇ:̌,UŞ)<ɱ͘ȋŔŉ͙YÔ2ʉʪ̫̳͘ȋŔ͙YO6˖̸˃ȻƖ (complex thalloid):RAVUÄð
!áİɒ9U r~khʕ@jvkhʕ=ǿ9èŦ+2͚̱¡Ǳȹ?§9OǒǪȂĔYO6è͌ʟ?
͜6:ʢSV8U͘ĩ͙͜r~khʕ?˵Ĉ?Ē)Yɧ-ˣƧ:+8͚b}pʃ?ĮŤ
SL6 UʪňPǱȹÄ?̛:ʢSVUûɢ!ȿɃ?r~khʕ?O?:À8U':!
ƪƳ)V8U (Wellman et al. 2003, Graham et al. 2004)23+͚@5"T:r~khʕ:è U
ǱȹÄ?ûɢ@§Ƀ·¸̭?ĮŤ S+ L6 58< (Walton 1925͙
ȿɃɮ?èŰtPèňʂʐ˞ǣ?ʎǥ@͚r~khʕ?ɨÚ!ĒɃ·?ʃ ͘Permian͙:§Ƀ·?
Ɍʃ ͘Triassic͙?ĸɖ͘P-Tĸɇ͢ʄ2Ø5ÿŸò͙?Įɀ˚ǹ?Ƀȹŀ̢ʑȩYɃ"ƀC͚0?Ƌ͚ȡǎû͚«ȶû+2ɂĸ=̘Ɣ+8͚̘ƔƷƹ+2':YɧĢ+8U (Wheeler 2000)
 r~khʕ@͚_pr~kh°ʕ:r~kh°ʕ=ŀí)VU (He-Nygren et al. 2006)_p
r~kh°ʕ@͚_pr~khɖ?L S<T͚_pr~kh (Blasia pusilla) :nfn
kh (Cavicularia densa) ? 2 ɮYěM˃ȻÄ?ʙ=@˃?R=O˙Uʉ éV̂L!
T͚â̛=ˊˋ͌YßɃ-Uɳ̳YO6è͌ʟ9̷͚ģźP̵ģźYO2<ʪňÄ@r~k
h°ʕ:ȈG8ŀİ9͚˃ȻÄ?Úɷ= 12 Î.6¶"͚ǧ@ƺ cm =¿CUʪňĥ?ľĺ͘˅ĺ͙! 2 ʉʪ¸¡?ćLYO6͚ʪňȇʉʪ!ȟƺè˓=Úɶ58 4 6=#CVU<;͚xkh°ʕ?ʪňÄ:ß̌-UȺƑYO6 (Duckett & Renzaglia 1993, Shimamura et al. 2005, ūǜS2006)_pr~kh°ʕ@͚r~khʕ?§9OǒOëǖ=èŦ+͚r~khʕ?< 9ĈŇ
ɒ<ƆƚɒȺƑYÔ2è͌ʟ:ʢSV8U (Forrest et al. 2006, Shimamura et al. 2012)ǂ͚
r~khYěMr~kh°ʕ͘ʄ 400 ɮ͙@͚ukkhɖ (Sphaerocarpales)͚oskhɖ
(Neohodgsoniales)͚cyer~khɖ (Lunulariales)͚r~khɖ (Marchantiales)

S<T͚ŀĀ?ɮ@r~khɖ=ěKVU (Long 2006)r~khɖ?¨˗<è͌ʟ?ʂʐ̬ËYĩ 2 =ɧ-r~khɖr~khɪ͘Marchantiaceae͙@͚r~khɖ?§9͚ǒOëǖ=èŦ+2è͌ʟ:ʢSV8Ur~khɖ@͚ȋŔŉ:ȋŔYèû-UĿŤǶ̍?˃ȻÄYO6':͚˃Ȼ
Ä?ʯ́=ʯ͕ȷ:¹ǯYèû+͚¹ǯ=@ʥć=RUǹǷ!UO?͘ǔʅ¹ǯ͙:<O?

M. Shimamura-2
BSJ- Revi ew 3: 85 ( 2012)

植物科学最前線 3: 86 ( 2012)͘ŷȪ¹ǯ͙?¥ǂ!èû-U͚ģȻ?ȴƖʻģ?§=ĿʉʪƖ?ȴƖʻY6#U':͚ɃȅģőPʪňÄY¶%U2N?Ⱥí<ɃȅǦ9U̵ģƢ̷͚ģƢYèû-U<;?ȖɃƆ˳9ȺƑ
7%SVU'VS?Ƕ̍@r~khYěMr~khɪ (Marchantiaceae) 9@ʷɒ3!͚0?
µ?ɪ9@'VS?Ɔ˳?̛͚U@ŀ̛è@̇û+͚țŃ+8U':OĿ(Boisselier-Dubayle et al. 2002͚ĩ 3͙
K2͚r~khɖ?ʪňÄ@͚µ?t^͌:ȈGƐŞ9Ƕ
̍!Ăʆ9͚˅ĺ! 1 ʉʪć͚ǧ@ɡ ǾŃ-Ur~khɪ9@͚Ŀƺ?Ş)<ʪňÄ!̷
ģƢ?¡̷̛͘ģź͙ Sƛį+8ɏɃ+͚c|͚Ñʺ˒͚ʽʰ:̝ÐÄɆǟ?ģő
9 3 ̠=Íˮ)V<!SƜ̨-Ur~khɪ@͚v`or~khţ͘Bucegia͙͚\cr~khţ (Preissia)͚r~khţ (Marchantia) S<Ur~khţ(Marchantia)@ʄ 40 ɮ!ɠSV͚ǅǚīâ=@͚M. polymorpha ssp. ruderalis͘r~kh͙͚M. polymorpha ssp. polymorpha͘vr~kh͙
͚Marchantia paleacea ssp. paleacea͘xr~kh͙͚Marchantia paleacea. ssp. diptera͘ t
r~kh͙͚Marchantia emarginata ssp. tosana͘|lr~kh͙͚Marchantia pinnata͘|{r~kh͙!èŰ-U(Bishler 1989, 1998, ȷǰĒǗ 2012)

ĩ̱͜¡Ǳȹ=%Ut^͌r~khʕ?ʂʐɒÁʞt^͌@ȿɃ?̱¡Ǳȹ?§9͚O5:Oëǖ=èŦ+2:
ʢSV8UʔɻǝǱȹ:jhǱȹ?ʂʐ̬Ë=68@Ǉɥ<ʎ˪!ç8<!͚ǒ̃?èǦʂʐōɒɣɲ?
Ŀ#@xkh͌:ʔɻǝǱȹ!ņŅʟ:<U':Yɧ+8Uĩ 2.r~khʕ?¨˗è͌ʟ?ʂʐ̬Ër~khYěMr~khɪ@͚r~kh°
ʕ?ɮ?ŀĀYĄNUr~khɖ?§9ǒ
Oëǖ=èŦ+2:ʢSVUShimamura et al. (2012)YO:=Æĩ

M. Shimamura-3
BSJ- Revi ew 3: 86 ( 2012)

植物科学最前線 3: 87 ( 2012)

ĩ 3.r~khʕ?¨˗è͌ʟ?ȺƑa. _pr~khɖ_pr~khɪ͚nfnkh (Cavicularia densa) ?ȴƖ
ʻģY6%2˃ȻÄ_pr~khɪ@͚ȋŔYO2<!˃ȻÄ?â̛=ˊˋ͌YßɃ)/UʭƝ!U˃ȻÄ?D4!ʉ #éV̂M̵ģź:̷ģź!<#͚̍ʀģ@˃ȻÄ=ƹĭ+͚ʪňÄ@˃ȻÄ?Úɷ=¶#b. cy
er~khɖcyer~khɪ͚cyer~kh (Lunularia cruciata) ?̷ģƢcyer~khɪ@ 1 Ť?ȋŔYO6̵ģź@ȴǧ̷ģź=Ñʺ˒!<ʪňÄ?ǧ!R#ɏ̕-U ǅǓİ?ȴƖʻģYO6c. ukkhɖukkhɪ͚eukkh (Sphaerocarpos donnelli)ukkhɪ@ȋŔYO2<ʉ˃ȻÄ?ʙ!˃
?R=éV̂L͚ʿ˃Ä=̃ľƆYO6 pjİ?̵øʰ:Ñʺ˒YŗɃ-Ud. r~khɖr~khɪ͚\
cr~kh (Preissia quadrata) ?̷ģƢ͘ŹŨŀōȷǰɠ©ȊƴƇ͙r~khɪ@ 1 Ť?ȋŔYO6̵ģƢ̷͚ģ
ƢYO6̷ģź@Ñʺ˒:ʽʰYO6e. r~khɖokhɪ͚okh (Conocephalum conicum) ?̵ģźo
khɪ@ 1 Ť?ȋŔYO6̵ģź@ȴǧ̷ģƢǧ@ʪňƹŰ?̴=ƕȮ=¿̨-UÑʺ˒!<ȴƖʻģ!<f. r~khɖodlkhɪ͚odlkh (Reboulia hemisphaerica ssp. orientalis) ?̷ģƢ:̵ģźodlkhɪ@ȋŔ! 13 Ť=ɏ̕-UȋŔ=ėûʁ!<®ČèǦ3%9<#͚ǱȹÄ?ʯ́ S?´ĭɒ<èǦOˏȴƖʻģ!<g. r~khɖov_khɪ͚xdthov_kh (Sauteria yatsuensis)?̷ģƢ͘ȷǰɠ©
ȊƴƇ͙ov_khɪ?ȋŔ@ 23 ŤȋŔ=ėûʁ!<̷ģƢ@˃ȻÄ?̋§=6#̵ģƢ@< ͚ȴǧȋŔŉ?̪đ̛!ǉƆȴƖʻģ!<h. r~khɖ_ekhɪ͚_ekh (Riccia fluitans)_ekhɪ@ȋŔ! 23 Ť=ɏ̕-U̵ģź:̷ģź!<#͚̍Ćģ:̍ʀģ@˃ȻÄâ̛=ȎɃʪňÄ=ǧ!<Ȍ§=Ș̑
+8Ƀʦ-UɮOUi. r~khɖhr~khɪ͚hr~kh (Dumortiera hirsuta) ?̵ģƢhr~khɪ@ȋŔPʯ͕ȷ!ɍ˸ɒ̵ģƢ̷͚ģƢYO6!̵ģƢǧ@̀ų=ɡȴƖʻģ!<

M. Shimamura-4
BSJ- Revi ew 3: 87 ( 2012)

植物科学最前線 3: 88 ( 2012)C    =./0%

 r~kh?è͌ōɒɣɲ@͚Marchantia polymorpha L.!1753Ÿ=Linnaeus=R58ˢ˾)V2':=̗U (Linnaeus1753)M. polymorpha L.͘Źʠ?r~kh͙=@ɃʦĮPɃʦɂĸ=R58͚
Ɔƚ=ĿǷƖ!U':!͚Linnaeusʳ˻P0?Ƌ?ɣɲ9˦NSV8T͚ƺĿ#?ɮâè͌
ʟ͘°ɮPĽɮ͙!ˢ˾)V8"2Burgeff (1943)@0VSYM. polymorpha (Ƚʠ)͚M. aquatica͚
M. alpestris?36?ȼɶ+2ɮ:+8ƻɁ+2͛0?Ƌ͚Burgeff (1943)?L:N23è͌ʟ@͚Ŀ#?ɣɲʣ=M. polymorpha?ɮâè͌ʟ:+8Đ%ÜVSV (Schuster 1992)͚ȿĭ@͞6?°ɮ
?LY˦NU?!ʷɒ:<58U (Bischler & Boisselier-Dubayle 1991, Perold 1999, Paton 1999)
è͌ōɒŒʠ!Ǐǋ952͚M. polymorpha L. (Linnaeus 1753)=̬+8@͚Bischler & Boisselier-Dubayle (1991)=RT͚Ĉˢ˾9ƂɅ)V8Uĩȸ (Dillenius 1741−1742)!f|t^:+8ƪŒ)V20?ƆƚɒȺƑ@͚ƍǟ͚M. aquatica:)V82Ǳȹ=ʴ-U͛ ȿĭ˦NSV8U͞6?°ɮ?4M. polymorpha ssp. polymorpha͘ƍǟ͚M. aquatica:)V
82O?͙@͚˃ȻÄ§ł̛=ȋŔ!èû/.͚ʯ́Ïʌʜ?ʹ!̉%8˙U2N=§ł̛?͗ʗ!̎ʓɒ9Ǉɞ9U͛)S=ʯ͕ȷ?¶ţȹ!Ýʙ9U':͚ɃȅģőPȴƖʻģ
YH:Z;6%<':<;9ȺƑ7%SVU¨=ťĮ?Ȍ́=èŰ+͚²ȳɒ<ɂĸ=@Ƀ
ʦ+<ǚ°ɮ@ǅǚ=OèŰ+͘üŬ 1987͙
͚vr~kh:RAV8Uĩ4=ŹŨŀōǱȹǸǚžƝˈ? Ÿ=ʟ͐əšȰhĈ9Ư̶)V2ǸǚYɧ-M. polymorpha ssp.
montivagans (ƍǟ͚M. alpestris:)V82O?)@͚˃ȻÄ?ŵ!2 cm=̕-U':OUŀİ?Ǳȹ9͚˃ȻÄ?§ł̛?͗ʗ!LSV<w9@͚ťĮY§ƒ=è?
˯Ř<ɂĸ=Ƀʦ+͘șŧ̃#?ķƖȤĮ9?Ư̶ˢ̦OU͙
͚²ȳɒ<ɂĸ=@Ƀʦ+<
ǚ°ɮ@'VK9?Ɲ͚ǅǚ9@èŰ!ɥ˦)V8<M. polymorpha ssp.ruderalis ͘ƍǟ͚
Ƚʠ?M. polymorpha:)V82O?͙@͚˃ȻÄ?§ł̛?͗ʗ͘ȋŔ!ɏ̕+<̛è͙!£̎ʓ9͚ʯ͕ȷ?¶ţȹ=̤ȁ!U':͚ȴƖʻģPǔƖɃȅģőYǆɔ=6%U<;?
ȺƑ!U²ȳɒ<ɂĸY§ƒ=üĀɀÝı=Ź#èŰ+͚āĀɀ=O̜ư̊Y§ƒ=ɬÜ!
LSVUǅǚ9O²ŕ?Ğ́9Ǎ̌=Ƀʦ+͚“r~kh”?ġĘ9ĠAV͚Ŀ#?ɣɲŔ9ɣɲǛƾ:+8ɅSV8U?@'?°ɮ9U Burgeff (1943) ?±̸œ͒@¡̅?3°ɮ?̙½ɒ̬ËYʢU¡9͚ʵğȝʎǥYɧ+8
U0V=RU:vr~kh:M. alpestris (ȿĭ͚ssp. montivagans:)V8UO?)?̫9@͚
H:Z;±̸!˵'S<!͚(#ű=Ƀ,2̸ɮ=r~kh=À2ƆƚYO6O?!Uǂ9͚r~kh@͚vr~kh:M. alpestris0V1V:?̫9Ŗǈ=±̸-U0VQ͚Ɖ@r~kh!͚vr~kh:M. alpestris?̸ɮ=˵Ĉ-U:ʢ2Schuster (1983, 1992)O0?ʢYƵƩ+͚r~kh?Ƀʦ!²ȳɒɂĸ=̮SVU': Sr~kh@͚ÚĔǌ·=Ƀ,2̸ɮ=˵Ĉ-Ù͚ų=ǁ+ɮ:ʢ20?ƋˏXV2\m^DNAĿİ?˞ǣY̌
,8͚ 6?°ɮ!̙½ɒ=Oýí9"U':@ǇS =<52!͚r~kh!µ?2°ɮ?̸ɮ
=˵ȥ-U:ˣƧ@ƎSV8< (Boisselier-Dubayle & Bischler 1989, Boisselier-Dubayle etal. 1995)ǨʹÄƺ@.V?°ɮO9ǚ (n=9̷̵͚?ƖǨʹÄYěM) 9U':!è 58
U (Bischler 1986).

M. Shimamura-5
BSJ- Revi ew 3: 88 ( 2012)

植物科学最前線 3: 89 ( 2012)ĩ ͛vr~kh(M. polymorpha ssp. polymorpha)?Ǹǚ ˃ȻÄ̷͘ǭ͙?ľƆ͛˃ȻÄ@ʉ̨#͚ŵ@  ͛èǦ?˝ż!r~kh:ȈG̣˝Ɯȵ+2̷ģƢ͘ɟą͙@˃ȻÄ?Ĳ̛̃#=Áʞ-U̷ģź@Ş)!͚ǧ@ :̨#¿CU'?Ǹǚ@ȴƖʻģY6%8<͛˃ȻÄ§ł̛¶̃͛§ł̛=ȋŔ!Ý#èû/.͚
§ł̛?͗ʗ!̎ʓɒ9Ǉɞ͘ɟą͙ʯ͕ȷ?¶ţȹ̤͛ȁ!<#͚Ş)<ʉʪ9ʙďSV͚Ýʙ͛͘ĩ  ?r~kh?ʯ͕ȷ?¶ţȹ:Ȉ˽/R͙Specim. exam. Japan. Pref. Gunma: Ozegahara, 1400 m alt. H. Ando 9143 (HIRO).

D

+&

 r~kh?ɃȕĔ?ǵɉYĩ 5 =ɧ-͛r~kh@µ?Ǳȹ:ė,#͚Ǯɘ?ĽûY¾¤·±·Yˏµ?jhǱȹ:ėǷ͚Ƀȕ?¨Ä:<UǱȹÄ͘˃ȻÄ͙@Ăɘ?̝ÐÄ9U
ǱȹÄ@̷̵ɋǭ9̵͚ǭ=@̍ʀģY6%U̵ģƢ̷͚ǭ=@̍ĆģY6%U̷ģƢ!ƆƜ)VUĿ#?t^͌9@͚Ƀȅģő!Ÿ̫?̮SV2ǌǖ?LƆƜ)VU!͚r~kh@Çľɒ
9̡͚ľ=8OåǖY̰8̨͚ǖ̫=X2Tǿ:̵ģƢ:̷ģƢ!ƆƜ)VU̡ľ?
ÎÄʟ9@ǊÚ=̵ģƢ!̷ģƢ=Úɶ58Ɯ̨-UÕę!U̵ǭ?̍ʀģ9ÆSV2ʀň!̷ǭ?̍Ćģ=ï̕+͚â̛9Đʀ!ˏXVU:͚ĐʀĆ@ɏɃYŇN͚˖ɘ?ʪňÄ!ƆƜ

M. Shimamura-6
BSJ- Revi ew 3: 89 ( 2012)

植物科学最前線 3: 90 ( 2012))VUʪňÄ@̷ǭ?̷ģź?¢̛=ƛį+8Ɯ̨-U͛
ʪňÄ?â̛9@ȟƺè˓!'T͚Ăɘ?ʪň!ƆƜ)VUʪňÄ SƷç)V2ʪň!ɏʻ+͚ĈʁÄ:ĠAVUÄðYʍ8Ǳ
ȹÄ!ƆƜ)VUǔƖɃȅY´+2ʪňƆƜ3%9<#͚ǱȹÄ?ȴƖʻģ?â̛=ƆƜ)VUȴƖʻ=R58Oǆɔ<ȴƖɃȅYˏ¸¢͚ɃȕĔ?ĕȆ̲9˙SVUƆƚ=68͚RTˤ+#˞˨-Uĩ 5͛r~kh?Ƀȕɂ

E33-4
 r~kh?ʪň@ 10–15 µm ɭż?ɀƆ9͚ʪňÄ SƷç)V2:͚̘ƅ<ɂĸǞº!ƻ
A»ɛ-U':<#͚234=ɏʻ-Uɏʻ@ÛÉŋɒ9͚ʪň!ÛĖƜYˏ':!̠˗
9U':!è 58U (Inoue 1960, Nakazato et al. 1999)ʪň@ɏʻ?̴=͚ŀŞ͝6?ʉʪ
YɃ,U£ɹè˓Yˏ͘ĩ 6a͙
'?4Ş)ʉʪ S@ǒë?¹ǯ͘ŷȪ¹ǯ͙!¿̨+͚
ŀ"ʉʪ@ƺżè˓-U':9͚ƺʉʪ!ê=̎<T͚Ǚɷ=¹ǯYO6ĈʁÄ:<U(Inoue 1960)͘ĩ 6b͙
0?Ƌ£˚ñ<ʉʪè˓Yˏ͚ĵȻ?ľƆYʍ8͘ĩ 6c͙͚Uɭż?
ŀ")=<U:͚͝ǂę=±¯=˚ñȀ+ʉʪè˓YʝT̄-ͅɷʉʪ!çȿ+͚ĈʁÄ¡=˃ȻÄǷ?ŷ2̛è!9"U (Leitgeb 1880)͘ĩ 6d͙)S=ĈʁÄ?ʉʪ!ƺþʉʪ=Ĺõ+͚ʧʯ́ǂę=Oè˓YˏǳƆ?ͅɷʉʪ!ƆƜ)VU:͚˃ȻÄ?ȣƯ̈͘wv͙!ƆƜ)V͚ćLYO52˃Ȼ?ǱȹÄ?ƆƜ!̪Ň-U (O'Hanlon 1926)͘ĩ 6e͙

M. Shimamura-7
BSJ- Revi ew 3: 90 ( 2012)

植物科学最前線 3: 91 ( 2012)ĩ 6͛r~kh?ĈʁÄa. ʪň?ǒë?ʉʪè˓9Ƀ,2Ş)ʉʪ S¹ǯ!¿̨-U͘ɟą͙b. ŷȪ¹ǯ͘ɟą͙YǙɷ=O6͚ƺʉʪ!̎<52ĈʁÄc. ëǖ?ĈʁÄ@ȺŒ?ʉʪè˓tYO2.͚Ƿ<ĵȻ?ľƆ
YO6d. ĈʁÄ¡=Ƀ,2ͅɷʉʪ͘ɟą͙
e. ͅɷʉʪ!Ƀ,U:͚0'!ȣƯ̈͘wv͙:<58͘ɟą͙͚ŷ́ɒ<˃ȻÄ?ƆƜ!ŇKUF$(>5)? Ɯȵ+2r~kh?ǱȹÄ͘˃ȻÄ͙@ŵ 8 mm

S 15 mm ɭż͚§ł̛?ćL@ 0.3 mm

S 0.6 mm ɭż9͚®ČȻ=èǦYʝT̄+8Ɯ̨-Ur~khɖ?˃ȻÄ?ǀ́@˖̸=èû+2ŤȻǶ̍:<58T͚ʌʜèû?Ăʆ<xkhʕ?˃ȻƖt^͌?Äð!Ăʆ˃ȻÄ (simple thalloid) :RAVU?=Ś+͚˖̸˃ȻÄ (complex thalloid) :RAVU͘ĩ 7͙˃ȻÄ?ͅɷ̛@ƒʲİ=ȣÜ+8T͚0?¶̃!ʉʪè˓?ɔZ<Ɯ̨Ȳ:<58UƜ̨ȲYǶƜ-Uʉʪʟ?§ł=@͚͜6?ͅɷʉʪ (apical cell) !L:NSVUͅɷʉʪ@Ğ́?
ʉʪ=ȈG8ŀİ9͚ʉʪ˳=ŘZ9T͚¦Ťè˓=R58Ƀ,2ʉʪ=R58Śɫ=ďTĨ
KV8U':9͚µ?ʉʪ:ýí9"Uͅɷʉʪ?Ğ́9@ȕɏ<ʉʪè˓!˵'58Uǂ9͚ͅɷʉʪʳÄ?è˓͊ż@Â#͚ʉʪYéTç-ǂęYĉŗ=ðƏ-U':9͚ǱȹÄ?ƆƚƆƜ=̠˗<ƈóYƦ58UjhǱȹ9@ͅɷʉʪ SéTç)V2͜6?ʉʪ͘qg|͙=Ɇǟ-UʉʪʟY [^|:RC͚
ǱȹÄYǶƜ-Uʌʜ!˚ñɒ=èû-U
jhǱȹ?ǱȹÄ@ [^|!ɰL̠<52Ƕ̍9T͚ [^|@ǱȹÄ?ʌʜèû

M. Shimamura-8
BSJ- Revi ew 3: 91 ( 2012)

植物科学最前線 3: 92 ( 2012)?ĲǚĂÁ:<58U:Uͅɷʉʪ?Ɔ͚ͅɷʉʪ?ʉʪè˓ǂę?ðƏ͚ [^|§9?ʉʪè˓t?˚ñƖ@͚ĕè͌ʟ?Ĳǚɒ<ƆƚƆƜYƵ̝+8Ur~kh?ͅɷʉʪ@ 4 6?ʉʪè˓́YO6ǳƆ9U͘ĩ 8͙ͅɷʉʪ?ʳɆ́͘ľɇ=ư-Ú͙
@Ǳȹ?ʯ́Ï=Õ8̝ʞ+8T͚ʯ́Ï Sɏ̕-UɿȜȉ:ʯ͕ȷ=R58Íˮ)V8
Uͅɷʉʪ SǱȹÄ?ʧ́Ï=éTç)V2qg|=Ɇǟ-Uʧ́ [^|͚ʯ́Ï=éTç)V2qg|=Ɇǟ-Uʯ́ [^|@͚˃ȻÄ?§ł̛YǶƜ-Uʌʜ
͘ːɓ:͓˳͙:<Uͅɷʉʪ?Ïǂ=éTç)V2qg|=Ɇǟ-UÏǂ [^|S@͚˃ȻÄYǶƜ-UÝ8?ʌʜ!èû-U͘ĩ 9, 10͙ [^|§9 4 Ħ?ʉʪè˓!'T͚Ïǂ [^|! 5 ʉʪ:<52ǌȲ9͚ʧ́ːɓʌʜ͘ːɓȋŔ͙͚ʧ͓́˳ʌ
ʜ͚ʡ̛͘˃ȻÄ?ʙ͙
͚ʯ́ːɓʌʜ͘ːɓʯ͕ȷ¹ǯ<;͙͚ʯ͓́˳ʌʜF:èû-U
ʉʪ!ȍŒ-U (Crandall-Stotler 1981)r~kh?ĴĖ͚Ïǂ [^|?ëǖ?è˓t
@͚ͅɷʉʪ:Ý#ė,9T͚®ǿͅɷʉʪ (secondary apical cell) :ORGUƭUʶYU2N͚ͅɷʉʪ:Ïǂ [^|?˫íYʌʜéȷ× Sìǀ-U?@̺+r~kh@Ă?ͅɷʉʪYO2<:˞̟+2̔Ċ?ɣɲÇOĿ#U+ +̎ʓéȷYÆ˕+͚ͅɷʉʪɗ¢?ʯ́ [^|!ʯ͕ȷ?ĈĲYƆƜ+<':YƪǸ=-VAͅɷʉʪYėŒ9"U ˃ȻÄ?ʧ́Ï@͚ÝÄ!fvf9˘XV2ːɓ!ɏ̕+8U0?ɗ¢=@͚ȋŔ(airchamber)!èû-UÎ?ȋŔ@ 6 ˝Ɔ?ŞŔYƆƜ+8U2N͚˃ȻÄYʧ́ SɜNU:͡˝Ɔ?ǹǷ!LUȋŔ@§ł=Áʞ-UȋŔŉ (air pore) =R58ľɇ:̌,8UȋŔŉ@͟ê?ʉʪ=ɂȻ=ďTĨKV2ʉʪ̫̳9T͚ʚǀ́9@Ǽİ=̝ʞ+2ʉʪʟ!
˜ř)VU͘ĩ 11͙ʷɒ=@r~khȋŔŉ@͚ʔɻǝǱȹ?ȋŉ?R<̪̩̒÷@+<:ˢ̅)V8U+ +r~kh͌?#6 ?ɮ͌9@ȋŔŉYĨMǼİ=̝ʞ+2ʉʪ?4͚ǒOŻ̛?ʉʪ!͚ʱĬ?Ľû=RTĽƆ+͚ȋŔŉ?̌ȋƖY˩ƻ+8U:ʢS
VUÇ!˜ř)V8U (Walker & Pennington 1939)ȋŔ?â̛=@˃ʖÄYĿ#ěMʉʪ͘ėûʁ; assimilatory filament͙!èû+͚ÛĖƜYƦ58U͘ĩ 11͙ȋŔŉ:ʉʪ̫̳YÔ2
ȋŔ@͚ȋŉYÔ2ʔɻǝǱȹ?˃ʤʌʜ:ėǷ=͚ÛĖƜʌʜY«ȶ SŎT66͚ƨƹ+P-ȋɘ?®̞ûȱʇYǱȹÄâ=ŝ#':=ƈɶ58U(Meyer et al. 2008)ȋŔŉ?̪̩
̒÷!< ͚£ŐÝ<O?9U:-VA͚ʔɻǝǱȹ?ȋŉ:ȈG8ˆƹ=RUȌ?ƲŃ!
ŀ":­Ȣ)VU+ +͚'?ˆƹÆɅ!͚Į́=ư+2˃ȻÄ¢́ S?Ȍ?ĜĎYŝ
8U? O+V<r~kh͌?ȋŔ?ɏɃ˵ȥ=̬+8@͚ʧ́?ǒľŤ?ʉʪê: 2 Ɋɖ?ʉʪê?̫=Ƀ,2ʉʪ̫̳=˵ȥ-U˨͘âɃ˵ȥ; Hofmeister 1851 <;͙:ʧ́ǒľŤ?
ʉʪėļ?ʉʪ̫̳=˵ȥ-U˨͘ľɃ˵ȥ; Leitgeb1880 <;͙?͝6!52!͚ tr
~khYɅ2 Apostolakos & Galatis (1982) ?ˤʉ<ʌʜōɒɣɲ@͚ȋŔ!ǇS =ľɃ˵ȥ
9U':Yɧ+8Ur~kh=8O͚ǱȹÄ?ͅɷ¶̃?ʌʜéȷY˜ř+͚ȋŔ?
ƨŀ̔ɭY̆:͚ǒľŤ?ːɓʉʪėļ=̳̫!Ƀ,͚0V!ƨŀ+8ȋŔ:<58UǷň!˜ř9"U͘ĩ 12a–c͙ ȋŔ?¢̛?˃ȻÄ?͓˳(ǩʌʜ; parenchymatous tissue)@˱ˈʌʜ:<58Uʉʪ@µ?
ʌʜ=ȈGŀ"#͚ʉʪ̫̳!ɏ̕/.͚˃ʖÄ@µ?ʌʜRTOŞ)#͚ȭɽɾ!ɏ̕+8

M. Shimamura-9
BSJ- Revi ew 3: 92 ( 2012)

植物科学最前線 3: 93 ( 2012)

U˱ˈʌʜ?ʉʪĺ=@ʉʪ?̨˼ǂę=ɗ±-Uǂę=̉Ǉ<ʥć!LSVU(ĩ 13)â̛=ŀ"<ȐÄ (oil body) YěMʉʪ͘ȐÄʉʪ; idioblast, oil cell͙@͚˃ȻÄ?ĕʌʜ9Ɲ=
èû-UĂʰʂbd9UȐÄ@͚^s^}ûĖȹ?ĖƜ͚ˇɰ?Ĵ:<58
U͘Suire et al. 2000͙ ˃ȻÄ?ʯ́Ï͘Į́=ư-UÏ͙=@͚ʯ͕ȷ (ventral scale) :¹ǯ (rhizoid) !èû-U(ĩ13)¹ǯ@˃ȻÄʯ́?ːɓʉʪ!è˓Y¾X.¿̨+8Ƀ,2O?9͚͜ʉʪ S<U¹ǯ=@͚ƽȲȻ͚ʗȻ?âɃʥć=RUʅǷYO6ǔʅ¹ǯ (pegged rhizoid K2@ tuberculate
rhizoid) :͚ƐŞ<ƽȲȻ?ʥć+ O2<ŷȪ¹ǯ (smooth rhizoid) ?͝6?t^!U¥ʣ@ɋ<UǽʬYƦ58T͚ɏɃĴƝP¿̨ǂęOɋ<58U (Kammerling 1897,
Schiffner1909, üŬ 1977)ǔʅ¹ǯ@Ł) 10-20 µm ɭż?O?!Ŀ#͚ʯ́ÝÄ SɃ,8͚˃ȻÄ?§ł̛Yɖƪ-R=ʯ́=ȑ58¿C8U(ĩ 14a, b)ǔʅ¹ǯ@͚ʥćYƆƜ-U̔ɭ9ʉʪ˳YŃȃʉʪ:<U!͚ǝû-U':9͚0V1V?̫=Ö#ȉʉɻȿ˰YîɅ+͚˃ȻÄʯ́ÝÄ=ȌYÈʏ-Uľ̛̌ŝ?ǽʬYƦ58U (McConaha 1941)ǂ͚ŷȪ
¹ǯ@͚ǔʅ¹ǯRTŁ#͘20–40 µm͙͚¨=§ł?͝ê?ʯ͕ȷP0?Ğ́ SɃ,͚ʉʪ˳
=ŘZ9U23+ʉʪĺ!ˉ2N͚˃ȻÄ?Ē̛è9@6EV2ƆȻ:<58U':
OĿŷȪ¹ǯ@͚¢ǂ=¿̨+͚˃ȻÄYĲȹ=ĪŒ+͚Ĳȹ SȌèP͏èYŝÜ-Uʍ
˹:O<58U͘ĩ 14c͙
r~kh͌?Ŀ#?ɮ9ˁǯYƆƜ-Uˁ͌!Ĳȹ=?CUŷȪ¹
ǯY̌,8͚ǱȹÄâ=ÊÜ-U':!è 58Uò̅+2Źʠr~kh? 3 °ɮ?4͚
M. polymorpha ssp. montivagans ?ǱȹÄ=@͚µ?r~kh͌:ėǷ͚ßɃˁ!LSVU!͚r~kh:vr~kh=@LSV< (Ligrone et al. 2007)ɵʇè?ĿĴƝ=ńZ9Ƀʦ-Uè͌ʟ9@ˁ͌:?ßɃ!Ɠ˗< ͚ˁ͌:?ßɃ=̬-UǽʬYŃ58+K52? O+V<r~kh?ǱȹÄː́=@͚vfz_!ɃƗ+8T͚Ǳȹ?Ƀ̨YÌ̏+8
U:ĳĝOU (Kutschera et al. 2007)
 ʯ͕ȷ@͚ʯ́ÝÄY˘R=ɏ̕-U͜ʉʪć?͕ȷȻ?ʌʜ9͚0?ƆP̝ê@͚r~kh͌?ĕè͌ʟ9ȺƑ!T͚ɮYėŒ-U̴? 1 6?ɖŏ:<Ur~kh9@˃ȻÄ?§
łYƬZ9͞ê.6͚ˠ͡ê=̝ʞ+͘ĩ 15a, b͙͚§ł?͝ê=@µ?͟ê=@<ɴ˵Ȼ?¶ţȹ!U͘ĩ 15c͙'?¶ţȹ@͚O:O:˃ȻÄ?ͅɷ¶̃9Ɯ̨ȲYʯ́Ï Sʧ́Ï=˘R=ɏ̕+8Íˮ+8U¶ţȹ@͚˃ȻÄ!Ɯ̨+ʯ͕ȷ?¶ɝÁʞ!Ɯ̨Ȳ S̹
VU=ƍ͚ʯ́Ï=Ƃ"̂KV8ʯ͕ȷ?òǂĞʙ̛?ɴ˵:+8ȄU͘ĩ 15d͙§ł?͝ê
?ʯ͕ȷ@˃ȻÄ?§ł̛=ȑ58̨#¢į-ƯèYO4͚ŷȪ¹ǯY¥Ï Sǝ>U͂Ȼ?
Ƕ̍YƆƜ+8T͚öȾ?ʸȌè̀̈?̌˹:<58U (McConaha 1941, üŬ 1977)͘ĩ 14b͙ǅǚɄr~khţ?µ?ɮ͌9@ʯ͕ȷ@ 4 ê=¦C͚r~kh?R=˃ȻÄ?ʙ Sŀ"#
ľ=@Lç-':@<2N̡͚ľ9ɮYìí-U2N?ƪǸ=<U͘ĩ 15d͙

M. Shimamura-10
BSJ- Revi ew 3: 93 ( 2012)

植物科学最前線 3: 94 ( 2012)ĩ 7͛r~kh˃ȻÄ?Ǻǀ́?Ƕ̍ʧ́Ï?ːɓʌʜ=@ȋŔŉ9ľɇ=̌,2
ȋŔ!èû+͚0?¢=èćǩʌʜ͓͘
˳͙!èû-Uʯ́ːɓ Sʯ͕ȷ:¹
ǯ!èû-Uǔʅ¹ǯ!ʯ͕ȷ=ȑR
=˃ȻÄ?ʯ́ÝÄ=èŰ-UŷȪ¹
ǯ@Ĳȹ=ę 58¿C8U˃ȻÄ?
Ɲ=ȐÄʉʪ!èû-U

ĩ 8. ͅɷʉʪ͘ǉą͙YěMʌʜéȷa. ˃ȻÄ?ʧʯ́=ŷˏ<éȷb. ʧʯ́=įɗ 6Ɯ̨˼=ŷˏ<éȷc. ʧʯ́=įɗ 6Ɯ̨˼YǺǀ-Uéȷd. ͅɷʉʪ:Ğ́? [^|?è˓t?ǹƁĩ

M. Shimamura-11
BSJ- Revi ew 3: 94 ( 2012)

植物科学最前線 3: 95 ( 2012)ĩ 9͛t^͌?è͌ʟ(:? [^|?è˓t:͚ʌʜ?ɏɃ˵ȥ:<Uʉʪ?̖a. jvkhʕj
vkh͘ʿ˃Ɩ͙
͜6? [^| S͜Ǥ?˃!ƆƜ)VUb͛jvkhʕ|^kh͘ùú-Uʿ˃Ɩ͙
͜6? [^| Sŀ"<Ï˃:Ş)<ʧ˃!ƆƜ)VUc.xkh°ʕxkhɖ͘ʿ˃Ɩ͙
͜6? [^| S͝6=éV̂Z3͚U@͝6=ƤTɌKV2͜Ǥ?˃!ƆƜ)VUd. r~kh°ʕ͘˃ȻƖ͙jvkhʕ:ė,è˓tYɧ-!͚jvkh9˃YƆƜ-Uʉʪ!͚˃ȻÄ?ʙ͘ʡ̛͙:<Uĩ r~khÏǂ [^|?ʚǀéȷʡ̛?ƟƆ?
ʉʪ

SéTç)V2ʉʪ?

ʌʜèû̔ɭYʹè%+8ɧ- ?ʉʪʟ@0V1V͚
ʡ̛?ƟƆ?ʉʪ

SéTç)V2͜ʉʪ=Ɇǟ-U

M. Shimamura-12
BSJ- Revi ew 3: 95 ( 2012)

植物科学最前線 3: 96 ( 2012)ĩ r~kh?ȋŔ:ȋŔŉ ˃ȻÄʧ́ SL2ȋŔŉ͠͡˝Ɔ?Î?ȋŔ?§ł̛=ȋŔŉ!̪đ-U͘ɟą͙˃ȻÄ§ł̛=@ȋŔ!èû+<͇ı͘ɟ͈͙!£̎ʓ=èŰ-UȋŔŉ?ǀ́ȋŔŉY:TK#
Ǽİ=̝ʞ+2ʉʪ@͚ȋŔŉ=́-UÏ!ʥć-UȋŔâ̛=@˃ʖÄ=ŘZ3ėûʁ!UːɓŤ=@˃ʖÄ!èû+<ʧ́Ï S˙2ȋŔŉ?ǒ¡̛˜ř?2N=ėûʁY̰Ċ+2ȋŔŉ͘ǉą͙@ãƆ?̪đ̛Y
O6ȋŔŉ?Ż̛ʥć+2  6?ʉʪ!/Tç+þŊƆ=̪đ+8Uĩ ȋŔ?ƆƜ̔ɭ ʾːɓʌʜ?ǀ́ȋŔŉ?˵ȥ:<Uːɓʉʪ̫=Ƀ,U-"̫͘ɟą͙:ƨŀ+66UȋŔ͘ɟ͈͙
ɏ̕§?ȋŔǒë=-"̫!Ƀ,2Áʞ͘ɟą͙9¡¢ǂę=ʉʪè˓!'U':9͚ȋŔŉĞ́?Ǽİ?ʉʪ̝ʞ!6#SVU͘ɟ͈͙ėûʁYÔ2ȋŔĩ ǩʌʜ͓͘˳͙?ʉʪ ǩʌʜ?˃ʖÄ@ėûʁ?O?:ȈG8Ş)#͚{ɾ!ɏ̕+8U§ł
?ʉʪ@ȐÄʉʪǩʌʜ?ʉʪ=@̉Ǉ<ŲȻ?ʥć!U

M. Shimamura-13
BSJ- Revi ew 3: 96 ( 2012)

植物科学最前線 3: 97 ( 2012)ĩ 14. ǔʅ¹ǯ:ŷȪ¹ǯa. ˃ȻÄ?ʯ́=ȑ58¿CUǔʅ¹ǯƽȲȻ͚ɂȻ?ʥćYO6b. ˃ȻÄʯ́§ł?ǔʅ¹ǯ?ǝ¥Ï Sʯ͕ȷ=˘XV8Uc. Ĳȹ=ę 58¿CUŷȪ¹ǯĩ 15. ʯ͕ȷ?Ɔƚa. ˃ȻÄ?ʯ́ʯ͕ȷ͘ɟ͈͙!ʯ́ÝÄY˘R=èŰ-UâÏ?ʯ͕ȷ
=@µ?ʯ͕ȷ=@<¶ţȹ͘ɟą͙!Ub. ʯ͕ȷ?èŰ?ǹƁĩ¥Ï= 3 ê.6͚Ėˠ 6 ê=̝ê-Uc. ǒâê?ʯ͕ȷ?¶ţȹĞʙ=̤ȁ!U͘ɟ͈͙d. ʧ́ S˙2˃ȻÄͅɷ̛ͅɷʉ
ʪ!Uwv?̛è@ʯ́Ï S/Tç+2ʯ͕ȷ9Íˮ)V8U͘ɟą͙ǒľê?ʯ͕ȷ@˃ȻÄ?ʙ S@Lç+8U͘ɟ͈͙G'4'4 ˃ȻÄ?ʧ́§ł=̛=ȑ58͚ȴƖʻģ͘ǠȻÄ; gemma cup͙!ƆƜ)VU͘ĩ 15a͙ȴƖʻģ?ʙ=éV̂L!T͚ľÏ=ª͈Ȼ?ɴ˵!U':@͚|lr~kh<;ŷȪ<ȴƖʻ

M. Shimamura-14
BSJ- Revi ew 3: 97 ( 2012)

植物科学最前線 3: 98 ( 2012)ģYO6̃ʙɮ:?˫íȲ? 1 6=<U͘ĩ 16a, b͙ȴƖʻģ?â̛9@͚Ż̛?ʉʪY˵ȥ:+8͚ȴƖʻ!ɶ4¡!UƆ9ǿ:ƆƜ)VU͘ĩ 16c͙ȴƖʻ@Ăʉʪ?ǧ:͝6?ȣƯ̈
YO4͚§ł̛=ćL?U͚ãƆ?ŷ2ƆȻY+8U͘ĩ 16d͙ȴƖʻ!Ɯ̨-U̔ɭ9͚ȣƯ̈YÍˮ-UɿȜȉ!ÆSVUȐÄʉʪP¹ǯ?ĈĲ:<UʉʪOèû-U!͚ȴƖʻģ?â̛=ɈK58U4@ȋŔPʯ͕ȷ@èû/.͚Ɔƚɒ=ʧʯƖYƩ2<ʉʪè˓@Ğʙ̛ÝÄ9ė˩ɒ='U2N͚ȴƖʻ@˃ȻÄ?R<ǂęƖ?U¿̨Ɯ̨Y/.͚ãƆ?ľƆYÍ52KKƜ̨-Uǿ:ÆSVƜ̨-UȴƖʻ@͚¯=ƥ+Ė':9ǧ?̛è!è̹+͚ȴƖʻģ?â̛=ˇɰ-Uˇɰ+2ȴƖʻ@͚ȴƖʻģ?ʙ!ƿN¡ǂ=Ź!52Ƕ̍ (splash cup)=<58U2N͚ȴƖʻģâ=˂42̻ɾ=R58͚@>:A)V8Ğ́=͎CƹUȴƖʻ!ȴƖʻģ S 120 cm ?˷̹=K9˺Z3:ˢ̦OU (Equihua 1987)ȴƖʻ!Į́=˂4U:͚Į́=ư-UÏ=¹ǯ!èû+͚ȣƯ̈?ͅɷʉʪ!ȕ÷YŇN͚ʧʯ
Ɩ?Uʌʜèû!ŇKUĩ 16. ȴƖʻģ:ȴƖʻa. ȴƖʻģb. ȴƖʻģ?ʙʙ:0?ľÏ=ɴ˵!Ƀ,Uc. ȴƖʻģ?Ż̛?ƆƜ̋¡?ȴƖʻȴƖʻ@ĂʉʪƖ?ǧ͘ɟą͙YO6d. ǧ Sè̹+2ȴƖʻǧ!è̹+2ɍ͘ɟ͈͙:͝6?wv͘ɟą͙H:!;!
 ̵ǭ:̷ǭ@0V1V̵͚ģƢ (antheridiophore)̷͚ģƢ (archegoniophore) :RAVU͚ÓȻ?ɃȅǦYèû-U͘ĩ 17, 18͙̵ģƢ?¡̛?ãɕȻ?Ƕ̍9U̵ģź (antheridial receptacle)͚
=@̍ʀģ (antheridium) !èû-U̵ģź@˃ȻÄ=R#À2Ƕ̍YƩ58T͚QUP =M. Shimamura-15
BSJ- Revi ew 3: 98 ( 2012)

植物科学最前線 3: 99 ( 2012)

8 ˓-UO?!Ŀ͘ĩ 17 a, c͙̵ģź!Ɯ̨-U̔ɭ9͚ʡ̛ Sʧ́Ï=ȋŔP̍ʀģ͚ʯ́Ï=ʯ͕ȷ:¹ǯ!èû-U͘ĩ 17b͙̵ģƢ?ǧ?ǀ́Y˜ř-U:͚˃ȻÄʯ́ S̎ʓ
-Uǔʅ¹ǯ?ǝY 2 êÔ8U':!è U͘ĩ 17c͙Ɯȵ+2̵ģź@͚ŵ?Źʡ̛Y
O62N̵͚ģź?¡̛=ȌY2NP-Ƕ̍:<58U͘ĩ 17b, d͙̷ģƢ¡̛?ÓȻ?̷
ģź (archegonial receptacle) @͚ȝ# 8-10 ˓+͚ƪȻɴ˵ (digitate ray) :RAVUǶ̍YO6͘ĩ
17a͙
Î?ƪȻɴ˵?̫=@͚ͅɷʉʪ!T̷͚ģź?Ɯ̨̔ɭ9̍Ćģ (archegonium) !
ǿ:ƆƜ)VUƪȻɴ˵?ɏ̕!ǘȵ<ʾ̷ģƢ9@͚̍Ćģ?̛͉!̷ģź?¡ǂPÏ
ǂ=ę8U!͚0?Ƌ̷͚ģź@ʌʜÝÄ!âǐ+<!SƜ̨-U2N͚̍Ćģ@˙ %¡͚ƛį+2Áʞ=ƆƜ)VUR=<U͘ĩ 18b)̷ģƢ?ǧ?ǀ́Y˜ř-U:̵͚ģƢ:ėǷ͚˃ȻÄʯ́ S̎ʓ-Uǔʅ¹ǯ?ǝY 2 êÔ8U (ĩ 18c)̷ģƢ?ǧ9@͚ȋŔPʯ͕ȷOèû-U̵ģƢ̷͚ģƢ?ǧ@Ɔƚɒ=@ʉ̨#ƌ̨+2˃ȻÄ:˞̟9"UK2̵͚
ģź̷͚ģźO¡̛=ȋŔ!èû+͚¢̛=@͕ȷǷ?Ƕ̍!èû-U<;͚Ĳǚɒ<Ƕ̍@˃ȻÄ:ė,9U̷ģź?ƪȻɴ˵?ǀ́Y˜ř-U:͚˃ȻÄ!âǐ+͚ɺȻ=<52O?
9U':!X U͘ĩ 18d͙

ĩ 17. ̵ģź?Ƕ̍a. ̵ģƢľƆb. ̵ģź?ʚǀ́c. ǧ?Ǻǀ́d. ̵ģź?ʯ́ʡ̛Y̰"͚ŀ̛è!ʯ
͕ȷ9˘XVU

M. Shimamura-16
BSJ- Revi ew 3: 99 ( 2012)

植物科学最前線 3: 100 ( 2012)ĩ 18. ̷ģƢ?Ƕ̍a. ̷ģƢľƆb. ̷ģƢ?ʚǀ́͘¢̛?ǧ?̃#͙
c. ǧ?Ǻǀ́d. ƪȻɴ˵?Ǻǀ́I818 ̍ʀģ@̵ģź?ʡ̛?ǳƆ?ʉʪ SéTç)V2ːɓ? 1 ʉʪ=˵Ĉ-U̍ʀģ!Ɯ̨-U̔ɭ9͚0VYďTĨMR=͚ȋŔ=R#À2͚ pjȻ?ʭƝ̍͘ʀģʭ͙!ƆƜ)VU͘ĩ 19a͙̍ʀģʭ?¡̛PĞ́=U̍ʀģYěK<ȋŔ:@ɋ<T͚̍ʀģʭ?â̛=@͚ėûʁ!èû/.͚ĂʉʪƖ?ɿȜȉ?L!şƺɃ,U͘ĩ 19b͙̍ʀģʭ?¡̛@͚Ş)ŉ9ľ̛:̌,8U̍ʀģ@ɡǧYO4͚1 ʉʪć?ohw|ʉʪ?â̛@ʀĈʌʜ (spermatogenous tissue, androgonial tissue) 9UʀĈʌʜ@Ŝǟʀň:<UʀĈʉʪ(spermatogenous cell) ?L S<UʀĈʉʪ@ʉʪ?¿̨Ɯ̨YH:Z;/.͚è˓ǂę!ɗ±-Uʉʪè˓YʝT̄-2N͚ʀĈʌʜ@Ŀƺ?Ş)<ǂƆ?ʉʪ S<U͘ĩ 19c͙̍ʀģâ
9ˏXVUǒƋ?ʉʪè˓9U͚ʀʉʪ (sperm cell) YƆƜ-Uʀȇʉʪ (sperm mother cell)?è˓?L͚Ǆŋ?ʉʪĺ=Ś+ʉʪĺ!ƿNǂę=ƮÜ)VU2N͚ʀʉʪ@ ˝Ɔ?ľƆY
O6͘ĩ 19d͙ t^͌?ǱȹÄ?Äʉʪè˓9@§ƒÄ?R<ɀȻ?ƐŞɻƆƜ§ƒ͘ǴƆƜÄ; polar

M. Shimamura-17
BSJ- Revi ew 3: 100 ( 2012)

植物科学最前線 3: 101 ( 2012)

organizer͙ SƐŞɻƆƜ!'U!͚0?â̛=@§ƒŞÄ!< (Brown & Lemmon 1990,
Shimamura et al. 2004)+ +͚ʀȇʉʪ?è˓9@͚1 Ś?§ƒŞÄ (centriole) YO52§ƒÄ(centrosome)!çȿ+͚ʈ̥Ä?ƐŞɻƆƜ§ƒ:<U (Carothers & Kreitner 1968)§ƒŞÄ@͚ʀʉʪ!ʀňFĽƚ-U̔ɭ9̓ȉ?Ĳ̛˔ʞ͘ĲŻŞÄ; basal body͙F:Ľû-U͘Moser &Kreitner 1970͙
ʀʉʪ!ʀňF:Ľƚ-U̔ɭ9͚ʉʪ˳?ʛŞ͚Ǯ?ĽƆ͚ʀňȺǔ?ƐŞɻǶ̍̓͘ȉ͚p^͚ĲŻŞÄ͙?ƆƜ!'U͘ĩ 20a–d͙ŐƜ+2ʀň@͚æ̶+2Ǯ
!ŀ̛èYĄNŲʫÄ̛YO4͚Ǯ=ȑR=p^:RAVUƐŞɻ?ǝ!U
̛͈?ʉʪ˳=@|j}\!T͚ĿŤǶ̍Ä=ư-U̓ȉĲ̛˔ʞ SƋǂ=ę%8 2
ǚ?̓ȉ!ŷˏ=¿C8Uš̛:̓ȉ?Ú=Oʉʪ˳?ĪKT!T͚š̛?ʉʪ˳=@˃ʖÄ:|j}\!U͘ĩ 20e, f͙jhǱȹ?ʀň?|j}\PʹʇÄ!Đʀ?̴
=Ćʉʪ=Ʃ4̂KVU':!˜ř)V8U (Yuasa 1952)+ +͚xkh͌:t^͌?ʀ
ʉʪ!ʀňF:Ľƚ-U̔ɭ9ʉʪ˳bd=Ɇǟ-U DNA ˍÛǨʹ?˿Ȳ!țŃ-U':(Izumi & Ono 1999, Shimamura et al. 1999)͚Ƿ<ɮ?̙½ɒ<˞ǣ9O˃ʖÄ:|j}
\!ȷ˛̙½-U':!ɧ)V8T (Pacak & Szweykowska-Kulińska 2003, McDaniel et al.
2007, Jankowiak-Siuda et al. 2008)͚µ?Ŀ#?̱¡Ǳȹ:ėǷ=͚jhǱȹ9Oʉʪ˳bd
!ȇƖ̙½-U2N?ǽǶ!ŋĭ-U:ʢSVUjhǱȹ?̓ȉ?ǀ́@͚Ŀ#?ɚǮ
Ƀȹ=ß̌? 9+2 Ƕ̍?ƐŞɻʟ̓͘ȉ˼ʁ͙ S<U!͚̓ȉ˼ʁ=u^~ľʮYO2<?!ȺƑ9Ur~khYěN͚jhǱȹ?ŐƜ+2ʀň͚U@Ľƚ̋§?ʀʉʪ=%UǶ̍?Ɔƚōɒɣɲ=@̉̔İ̼ň͋Ɛ̧YɅ2Ŀ#?ɣɲÇ!T͚p^YǶƜUƐŞɻ?ƺP͚ĲŻŞÄ?̝ʞ<;͚è͌ʟ(:=ȺƑɒ<Ɔ˳!˙3)V8U (Carothers& Duckett 1980, Renzaglia & Garbary 2001)
 ̍ĆģO̷͚ģź?ʡ̛?ːɓʉʪ=˵ȥ-U!̷͚ģź@͚ŷ́ɒ=Ɯ̨-U̵ģź:̖5
8͚˄+#âǐ+8Ɯ̨-U2N͚UɭżƜ̨+2̷ģź9@͚ǁ2<̍Ćģ@̷ģź?¢̛9ƆƜ)VU̍Ćģ@͜ʉʪ=˵ȥ+͚ʌʜ?âľYè%Uʉʪè˓=R58ǒë=â̛=Ƀ
,2ʉʪ!͚ĆĈʉʪ:<U͘ĩ 21a͙ĆĈʉʪ@̍Ćģ!Ɯ̨-U=ƍè˓+͚Ŝǟ̍Ćģ?̪đ̛:<UǬʉʪ (cover cell)͚4 Î¸¡?͉ȧʉʪ (neck canal cell)͚Ćʉʪ=ư-Uʯȧʉʪ͘ventral canal cell͙͚Ćʉʪ (egg) YƆƜ-U(ĩ 21b)Ǭʉʪ:͉ȧʉʪ!͚̍Ćģ?Ɯȵ̔ɭ9ŪĻ-U':9͚ʀň!ĆʉʪFï̕-U2N?̌˹!̪#͘ĩ 21c, d͙ʯȧʉʪ@Ş)#<U!͚ĐʀǌK9Ȅŋ-U':!Ur~khʕ9@̛͉YǶƜ-U͉ʉʪ!͡ê=¦C͘Ǻǀ́9͉ȧʉʪ?ĞTY 6 Î?ʉʪ!ďTŮ#͙
͚0?â̛!͚Đʀ?̴=ʀň!ĆF:ï̕-U
̌˹:<U͘ĩ 21d͙xkhʕ9@͉ʉʪ! 5 ê=¦E?!áİ9UŐƜ+2̍Ćģ@pjƆ9͚ʱSZ3ʯ̛ (venter) :̨̛͉ (neck) YO4͚ʯ̛?â̛=@͜Î?Ćʉʪ!U̍Ćģ?Ĳ̛@ȗãɺȻ?ʌʜ=ďTĨKV8U͘ĩ 21e͙'?ãɺȻ?ʌʜ@Đʀ!'U:͚̍ĆģYøL̂MˑȻ?ʌʜ=ɏ̕+͚Ñʺ˒(pseudoperianth):RAVU

M. Shimamura-18
BSJ- Revi ew 3: 101 ( 2012)

植物科学最前線 3: 102 ( 2012)ĩ  ͛̍ʀģ?Ƕ̍ ʾ̵ģƢ?ʡ̛?ʚǀ́̍ʀģ?ʚǀ́ʀĈʌʜǀ́ǂƆ?Ş)ʉʪ9ĄN
SVUʀĈʌʜǀ́͘ʀňĽƚǖ͙ʀʉʪ@ ˝Ɔ?ľƆYɧ-ĩ 20. ʀň?Ƕ̍:ƆƜ̔ɭa.ʀȇʉʪ?ƿNǂę?ʉʪè˓b. ̓ȉ͚p^ƐŞɻ?ƆƜc. ʉʪ˳?̇û:Ǯ?æ̶d. p^=ȑ52Ǯ?¿̨e. ʀň?ǹƁĩf. ƐèŶȞ͋Ɛ̧9˜ř+2r~khʀňa–e.
Carothers & Kreitner (1968), Shimamura et al. (1999)Yċʢ=Æĩ

M. Shimamura-19
BSJ- Revi ew 3: 102 ( 2012)

植物科学最前線 3: 103 ( 2012)ĩ 21. ̍Ćģ?ƆƜ̔ɭa. ʾ̍Ćģ:â̛?ĆĈʉʪ͘ɟ͈͙?ʚǀ́b. ̍Ćģ͘ĆʉʪƆƜɗƋ͙?ʚǀ́c. ̍Ćģʯ̛?ʚǀ́͘ʯȧʉʪ:͉ȧʉʪ?̇û͙
d. ̍Ćģ̛͉?Ǻǀ́͉đ͘ɟ͈͙@ 6 ʉʪê9ĨKVUe. ŐƜ+2̍Ćģ

A@1 ̵ģź=Ȍ!˟V8̵ģź?ĕʌʜ?ʉʪ!ʱƃ-U:͚0?Ĭô9̍ʀģ!ŪĻ+͚̍ʀģʭ?¡̛?ŉY̌,8ʀň!̵ģź¡=2K52Ȍ§FĤç-U͘ĩ 22a, b͙'?ŉ?ʚǀ́@\vİ9͚çđ!Ƚ#<58T͚Ǽİ?ʚǀ́Yɧ-̌ų?ȋŔŉ:@͚ƆȻ!ɋ<58
U͘ĩ 22c͙r~khɖ=@̵ģź=ǧ!<ɮOĿ!͚0VS@̵͚ģź?̍ʀģʭ?ŉS͚ɳȋ§=ʀňY 20 cm ̃#?͔)K9̽Ȼ=Ĥç+͚͍=RUʀň?ƹŰYˏ(Shimamura etal. 2008) ³?:'W͚'?R<ɳȋ§F?ʬ÷ɒ<ʀňĤç@r~kh9@ɠSV8<
r~kh@ʷɒ=@̵ģź?¡=ȦK52͚ʀňYěMȜȫ!̻Ȍ=R58@>:A)V͚Ğ
́=͎CƹU':9ʀň!ƹŰ+8U:ʢSV8UDuckett & Pressel (2009)!v
9ɝʹ+2ȜȫY̵ģƢ?¡=ȫ¢+2œ͒=RU:̵͚ģź?¡9˺>2Ȝȫ@ǒŀ 30
cm ?˷̹=͎Cƹ52ǂ9̵ģƢ SĜĎ)V2vȨȜ!͚͜ǌ̫¸â=͚ʄ
10 cm ?ŀ")?j~?ʯ́ÏÝÄ=͚ǔʅ¹ǯ?ǝ=ȑ58Ź!U':Oè 52K2͚
̷ǭYɝʹ+2Ȝȫ=Ț+2:'W͚3060 è9̷ģƢ?¡̛=K9ɝʹ+2Ȝ!ï̕+2'VS?ʎǥ@͚˃ȻÄ?ʯ́P̷ģƢ̵͚ģƢ=ŋĭ-Uǔʅ¹ǯ?ǝ!͚ǱȹÄÝÄ=ȌèY
ˏ"ȠS/U3%9<#͚Đʀ=8Oʀň?ɬ÷ʍ˹:+8ǽʬ+8UēʬƖYɧ+8

M. Shimamura-20
BSJ- Revi ew 3: 103 ( 2012)

植物科学最前線 3: 104 ( 2012)U̵ǭ?̃#9Ƀʦ-U̷ǭ?̷ģź9@͚)K*K<ɏɃȆ̲?ʪňÄ!LSV͚ǧ=̃
Ï=ǁ2=ÆSV2̍Ćģ9@͚Đʀ+2A T?O?OLSVU̍Ćģ@̷ģƢǧ!ɏ̕+8̷ģź!Ʃ4¡&SV2:Oʒʓɒ=ƆƜ)V͚Đʀ@̷ģź!Į¡ Sɶ4¡!52Ȼƚ
9Oʒʓɒ="8U':!ɧĢ)VU (Parihar 1956)ɸʣOœ̴=̷ģź?ǀ́Y˜ř+8
L2:'W̷͚ģƢ?ǧ?¹ǯǝ@̷ģź9Ǧè V+͚Î?ƪȻɴ˵â̛?¹ǯǝ:ʓ"=<58U':!è 52+2!58Î?ƪȻɴ˵?Ĳ̛?̫=Áʞ-U̍Ćģʟ@͚¥ÏY¹ǯǝ=ďTĨKV8U͘ĩ 23͙ʀň@̷ǭ?˃ȻÄ?ʙ=)ï̕-VA͚̍Ćģ?
(#̃ÒK9͚¹ǯǝ?ȉʉɻȿ˰=RTĐ÷ɒ=̒AVU?9@<3W ʀň!̍Ćģ
S?ÅS ?˧ħȹ˳=˧XV8Ȕ$Ɠ˗!U:-VA͚1 mm ¸¢?(#ɡ˷̹ O+V<ĩ 22. ʀň?Ʒça. ̵ģź?¡?Ȍ§=Ĥç+2ʀň(ɑȯ+8Ưè)b. ̍ʀģʭ?¡̛?ʀňĤçđ͘ɟą͙:Ğ́?ȋŔŉ͘ɟ͈͙̍ʀģʭ=@ėûʁ!ɏ̕+<2N͚¡̛ SLU:ȋŔ=ĨKV2͗5J͇ı:+8L:NSVUc. ʀň?Ĥçđ͘ɟą͙:ȋŔ?ʚǀ́͘ɟ͈͙
Ĥçđ?ǀ́@\vİ9͚Ǽİ?̪đ̛YO6ȋŔ
:@ƆȻ!ɋ<U

M. Shimamura-21
BSJ- Revi ew 3: 104 ( 2012)

植物科学最前線 3: 105 ( 2012)ĩ 23. ̷ģź?ƪȻɴ˵Ĳ̛?Ǻǀ́̷ģƢ?â̛=@˃ȻÄʯ́ Sʓ"=<52ǔʅ¹ǯ?ǝ!ÜT̂Z9
U̍Ćģʟ@ƪȻɴ˵F:ʓ#ǔʅ¹ǯ?ǝ=ďTĨKV8U':!è Uʀň!¹ǯǝYʍɆ+8ɬ÷-UĴĖ͚ƱŒ)VUĐʀ?ʍ˹Yɟą9ɧ-

AA3 Đʀ+2Ć@͚̍Ćģ?â̛9ʉʪè˓Y̪Ň-UK.͚¡¢ǂę=è˓+͚¡¢͝6?ʉ
ʪ!ƆƜ)VUʓ80V=ɗ±-UR<ę"9è˓!'U͘ĩ 24a͙̛͉=̃Ï?ʉʪʟ9͚¦Ťè˓!'T͚ľɇ=ư+<âÏ?ʉʪ!Ƀ,2:"=͘ĩ 24b͙͚ľÏY\
]zn_͘amphithecium)͚âÏYa}zn_ (endothecium) :RC͚t^͌9@\ ]zn_ S˅ĺ?L!Ƀ,͚a}zn_ S@ʪĈʌʜ͘ʪň:Ƅʁ͙?L!Ƀ,Ux
kh͌?ŀ̛èP͚q͌?̛9@a}zn_S͚˅?§ƒ̛?˼§͘j;columella͙:RAVUƵƩʌʜ?L!èû+͚ʪĈʌʜ@\ ]zn_=˵ȥ-U ò̅+2R=͚Đʀ!'52̍Ćģ?Ĳ̛ S@Î?̍ĆģYøL̂MR=͚Ñʺ˒(pseudoperianth)!ɏ̕-U)S=0?ľÏ S͚ʽʰ (̷ʽʰ; female involucre) 9Íˮ)VUʽʰ@͚Î?ƪȻɴ˵?Ĳ̛?̫=Áʞ-U̍ĆģʟYďTĨMR=ɏ̕-U (Parihar 1956)
̍Ćģ?ʯ̛@͚ʪňÄ?Ɯ̨=ė˩+8Ɯ̨+͚ʪňÄYɗưøMc| (calyptra) :<M. Shimamura-22
BSJ- Revi ew 3: 105 ( 2012)

植物科学最前線 3: 106 ( 2012)U6KT͚ʪňÄ@âÏ S͆=͚c|͚Ñʺ˒͚ʽʰ=RT 3 ̠=Íˮ)V8Ɯ̨U(ĩ 24c, d, 26a)c|:Ñʺ˒@͚Đʀ!'S<̮Tɏ̕+<ʿ:˃?èû!Ǉɞ<ʿ˃Ɩt^͌9@͚̍ĆģYÍˮ-U2N?˃ɒʌʜYʽ˃ (bract) :RC͚0?â̛=
U͚ĞĨ?˃!ɎĖ+8ʪňÄYøL̂Z3R=LUˑȻ?ÍˮʌʜYʺ˒ (perianth):RZ
9Ur~kh?R=͚˙ %¡˃!<˃ȻƖt^͌9@͚ėǷ?ʌʜ!ʿ˃Ɩt^
͌:@ɋ<Uɋ<UĘɫ9RAVU!͚ʽ˃:ʽʰ͚ʺ˒:Ñʺ˒@͚0V1Vɘė<ʌʜ:ʢSVU jhǱȹ9@̌ų͚ʪňÄ@¡ǂ=ę 58Ɯ̨-U!͚r~khɖ?Ŀ#?ɮ9@̍Ćģ!
̷ģź?¢̛=¢ę"=68U/9͚ʪňÄ@ƛį+͚¢ǂ=ę 58Ɯ̨-UʪňÄ@µ?t^͌:ėǷ͚˶͚ǧ͚˅͘ʪňĥ͙?͞6?̛è S9"8U͘ĩ 24d͙˶ (foot) @͚̝ÐÄ:?̎ʎ̛è9͚̝ÐÄ SʪňÄF?ǫ͏̀̈YƦ58U˶:̝ÐÄʌʜ!ư-ƯÁYʨŽ (placenta) :RC͚ʨŽ9@͚̝ÐÄ͚ʪňÄčǂ?Ï?ʉʪ!˖̸<ʥćYƆƜU'?ʥć@̝ÐÄ–ʪňÄ̫?ȹ˳̀̈:̬̎!U:ʢSV8Uǧ@˶:˅YưʓƯè9͚ʪňƹŰ?̴=¿̨Ɯ̨-U':9͚˅YÍˮģő?ľ=ƥ+ç-˅@͚͜ʉʪć?˅ĺ:0?â̛?ʪĈʌʜ (sporogenous tissue)

S<UʪĈʌʜ=@͚Ƅʁȇʉʪ (elatermother cell, elaterocyte) :ʪĈʉʪ͘sporogenous cell͙!Ś=<58èû-Ur~kh9@Ƅʁȇʉʪ@H:Z;ʉʪè˓YˏX.͚¿̨Ƀ̨?LYˏ͚Ƅʁʉʪ (elater cell) :<U͘okhɪ<;9@ƄʁȇʉʪOè˓+8Ĺõ-U͙ʪĈʉʪ@ 4 Ħ S 5 Ħ?ʉʪè˓Yˏ͚1 6?Ƅʁʉʪ=Ś+ 16−32 Î=Ĺõ+͚ȟƺè˓Yˏʪňȇʉʪ (spore mother cell, sporocyte):<U͘ĩ 24e͙r~khɖ9@͚r~khYěN8ȟƺè˓ǌ?bd?̝èPʉʪè˓˔ʞ?Ɔƚ!R#˩GSV8U(Brown et al. 2007, 2010, Shimamura et al. 2012)r~kh9@ʪňȇʉʪ@23͜6?ʹʇÄ͘˃ʖÄ͙YO4͘ĩ 25a͙͚Ǯ?è˓=Úɶ58͟6=è˓+͚Ŝǟ?
ʪň:<Uʉʪ˳͇ı=ɬ÷-UǮ?è˓@˃ʖÄ?̝è=R58S ,NȍŒ)V2ʉʪ
ǴƖ=ƍ58̏ˏ-U':!è 58U͘ĩ 25b͙t^͌9@˅?§ƒ̛=Ƅʁǝ:RAVU͚ʪňYèû/.Ƅʁ!ǝȻ=èû-U͇ı!ŋĭ-U':!U!͚r~kh?˅?â̛@ʪň:Ƅʁ!Ƿ=èŰ+͚Ƅʁǝ@˦NSV<͘ĩ 26a͙ʪň@͜6?ʪňÄ2T 300,000 Î¸¡͚͜6?̷ģź S 7,000,000 Î¸¡ɄɃ)VU:?ƱŒ!U (O’Hanlon 1926)͜6?̷
ģƢ@Ǌ

Sɩ=

%8ǒŀ 100 Îɭż?ʪňÄYʒʓɒ=ƆƜ-U:ĳĝ (Duckett &Pressel 2009) OU?9͚œ̴?͜6?̷ģƢ2T?ʪň?ɄɃ̢@O5:Ŀ O+V<ȟƺè˓ʋ¬Ƌ͚˅ĺ@ɂȻ?ʥćYƆƜ+͚ǒʋɒ=ʉʪ˳YŃ͘ĩ 26b͙Ƅʁʉʪ@ŭŮ"?ˎǃȻ?ʥćYƆƜ+͚ǒʋɒ=@ʉʪ˳YŃ͚ȃʉʪ9UƄʁ (elater) :<U͘ĩ 26c͙ʪňÄ@͚̍Ćģ=Ɇǟ-Uc|=øKV2Ȼƚ9Ɯ̨-U!͚ʪňYƹŰ-U̴=@͚ǧ!¿̨+͚˅@c|YɤT͚)S=c|Y˘Ñʺ˒:ʽʰY',̪%8͚ɳ§=̾ç-U͘ĩ 26d͙
0?Ƌ͚˅ĺ!ʪňÄ?Úɷ̛ S£˚ñ=ɤV͚Ƅʁ!«Ȥ̒÷-U':9ʪň!ɳ§=Ʒç)VU

M. Shimamura-23
BSJ- Revi ew 3: 106 ( 2012)

植物科学最前線 3: 107 ( 2012)ĩ 24. ʪňÄ?ɏɃ̔ɭa. ʩ͘4 ʉʪǖ͙?̉̔×ĐʀĆ?ǒë?è˓́͘ɟ͈͙=Ś+͚ɗ±-UʉʪǢ!ƆƜ
)VUb. a}zn_!Ƀ,2ʩĐʀĆ?ǒë?è˓́͘ɟ͈͙RT̛͉=̃Ï S˅!èû-Uc. ʩ?Ǻ
ǀ́ʩ@͚̍Ćģ?ʯ̛=Ɇǟ-Uc|͚̍Ćģ?Ĳ̛ Sɏ̕-UÑʺ˒=ďTĨKV8Ɯ̨-Ud. ˶͚ǧ͚ʪňĥ͘˅ĺ:ʪĈʌʜ͙?͞6?͇ı!èû+2ʪňÄ?ʚǀ́e. ȟƺè˓ɗò?ʪĈʌʜ?ʚǀ́Ƅʁʉʪ:ʪňȇʉʪʟ!±¯=̝ʞ-Uĩ 25. ȟƺè˓a. ʪňȇʉʪ?̉̔İ̼ň͋Ɛ̧=RU˜řP: ʹʇÄ͚N: Ǯb. ȟƺè˓ǌ?ʹʇÄ͘ʖ͙:Ǯ͙̿͘?ƫ÷ Ǯ?è˓=Úɶ4ʹʇÄ!è˓-UʹʇÄ?ː́=@-v͖͘ʹ͙!ŋĭ+͚ȟƺè˓
ǖ?ƐŞɻ͘˴͙?ƆƜ?˶Ĵ:<U

M. Shimamura-24
BSJ- Revi ew 3: 107 ( 2012)

植物科学最前線 3: 108 ( 2012)ĩ 26. ŐƜ+2ʪňÄ?Ɔƚa. ȟƺè˓!ʋ¬+2ʪňÄ?ʚǀ́b. ˅ĺ?ɂȻʥćc. ʪň:ƄʁƄʁ@ 2
̠ˎǃ?ʥćYƆƜ-Uˎǃ?ę"@Ý8ŭŮ"d. ǧ!¿̨+͚ʽʰ?ľ=̾ç+2˅AB

=. ./0#6< ǚɯ9ʊ´+2 M. polymorpha (Źʠ?r~kh)?è͌ōɒɣɲ@¨=w=èŰ-U
ǱȹYÙ=ˏXV8"2ʍʘ!T͚\o\Įı?Ǳȹ=68@͚ˤʉ<è͌ōɣɲ͚̙½ɒ˞ǣ@<)V8<ǅǚ9@͚üŬ (1987)!ʟ͐əšȰhĈ9Ư̶)V2ǸǚYO:=͚
vr~kh͘M. polymorpha var. aquatica :+8ėŒȿĭ? ssp. polymorpha =ɘƅ͙Yĳĝ+2':¸ľ=͚Źʠ?r~kh?ɮâè͌ʟ=̬-UɣɲÇ!<r~kh@è͌ōʣPƯ̶ŕ
=:58͚KT=O˻̃<͑O?9+ <2Nȓƙ!ơXV8'< 52:Ȳ@ĚN<)S=ǅǚ=@͚ǨʹÄ¡?ǮŞÄƆƜ̛Á (nucleolar organizer) ?Áʞ!wɄ
?ĕ°ɮ:@ɋ<UÎÄʟ!U':!ĳĝ)V8T (Bischler 1986)͚iǶ̍?̖!
U':!ɧĢ)V8U M. polymorpha ssp. montivagans =̬+8@͚w¸ľ9O͚\ c (Perold 1999)͚üɼ͘Schuster 1992͙
͚n\ (Konstantinova & Bakalin 2009)͚̈́ī (Choi et al. 2010)͚<;͔ʘżU
@͔Ǹ͔?ĮıY§ƒ=¤ɇĕĮ SɃʦ!ĳĝ)V8T͚ǅǚīâ9Oǁ2<˩ǪPǸǚ?äǲˡ!ǕKVU͞6?°ɮ̫?0V1V?§̫ɒ<Ɔƚ?Ǳȹ!ŋĭ-U':@͚è͌ōɒ<˞̟!ɣɲʣ=R58ɋ<UĈħ:O<58"2ÇAȤȬ<ɂĸ9Ƀʦ+8Ur~kh@vr~kh=À2ƆƚYɧ-':!U͘üŬ 1987, Schuster 1992͙
³Ƌ?˞ǣ9͞6?
°ɮP0?§̫İ?Ǳȹ?̙½ɒœÄ͚Ɔƚ?ēĶƖ?ŵ!ǇS =<U':YǖƊ+2

M. Shimamura-25
BSJ- Revi ew 3: 108 ( 2012)

植物科学最前線 3: 109 ( 2012)

 r~kh?ľ̛Ɔƚ=̬+8@ 200 Ÿ¸¡=X2Uɣɲ?ȂĔ?§9͚ˤʉ=˜řˢ̅)V
8"223+͚ȋŔ͚ʯ͕ȷ͚ǔʅ¹ǯ̷͚ģƢ̵͚ģƢ:52r~kh°ʕ=Ⱥǔ?ʌʜ?̏ûōɒ˵ȥ͚µ?jhǱȹ?ʌʜ:?ɘėƖ=68@͚'VK9˭˪!£˶+82ȿĭ͚̱¡Ǳȹ=%Ut^͌?ʂʐɒÁʞPjhǱȹĕè͌ʟ?â̛?ʂʐ̬Ë!+3=ǇS=<T͚Ɔƚ?̏û=̬XU˭˪Yʂʐ̬Ë=Ĳ78ˏUɂĸ!ƻ58"2r~khY{Ǳȹ:+2ɣɲ?̏Ţ9͚̱¡Ǳȹ=Ǎ̓ɒ<ȿ˰?Ɂ˞!ȝKU':?L<S.͚jhǱȹȺǔ?̏ûɏɃōɒɣɲ?ĲɦOɥɶ9"U?9@<3W 'VK9O5BS͚è͌ʟPɮY˦˫-U2N?
ƆƚƆ˳:+8?Lȓɖ)V82jhǱȹ?ľ̛Ɔƚ?ĿǷƖ=68͚àÄɒ<ʌʜ?ɘėƖY̙½ň
ɏȿ9ǲˣ+͚̏ûɏɃōɒ˜Ȳ9ɣɲY̏NU':!9"U O+V<,"*
Apostolakos, P., & Galatis, B. 1982. Studies on the development of the air pores and air chambers ofMarchantia paleacea. Protoplasma 128: 120-135.
Bischler, H. 1986. Marchantia polymorpha L. s. lat. karyotype analysis. J. Hattori Bot. Lab. 60: 105-117.
Bischler, H. 1989. Marchantia L. The Asiatic and Oceanic taxa. Bryophytorum Bibliotheca 38: 1-317.
Bischler, H. 1998. Systematics and evolution of the genera of the Marchantiales. Bryophytorum Bibliotheca
51: 1-201.
Bischler-Causse, H., & Boisselier-Dubayle, M. C. 1991. Lectotypification of Marchantia polymorpha L. J.
Bryol. 16: 361-365.
Boisselier-Dubayle, M. C., & Bischler, H. 1989. Electrophoretic studies in Marchantia polymorpha L. J.
Hattori Bot. Lab. 67: 297-312.
Boisselier-Dubayle, M. C., Jubier, M. F., Lejeune, B., & Bischler, H. 1995. Genetic variability in the threesubspecies of Marchantia polymorpha (Hepaticae): isozymes, RFLP and RAPD markers. Taxon 44:
363-376.
Boisselier-Dubayle, M. C., Lambourdiere, J., & Bischler, H. 2002. Molecular phylogenies support multiple
morphological reductions in the liverwort subclass Marchantiidae (Bryophyta). Mol. Phylogenet. Evol.
24: 66-77.
Brown, R. C., & Lemmon, B. E. 1990. Polar organizers mark division axis prior to preprophase bandformation in mitosis of the hepatic Reboulia hemisphaerica (Bryophyta). Protoplasma 156: 74-81
Brown R. C., Lemmon B. E., & Shimamura, M. 2007. Transformations of the pleiomorphic plant MTOC
during sporogenesis in the hepatic Marchantia polymorpha. J. Integr. Plant Biol. 49: 1244-1252.
Brown, R. C., Lemmon, B. E., & Shimamura, M. 2010. Diversity in meiotic spindle origin anddetermination of cytokinetic planes in sporogenesis of complex thalloid liverworts (Marchantiopsida) J.
Plant Res.123: 589-605
Burgeff, H. 1943. Genetische Studien an Marchantia. Gustav Fischer, Jena, Germany.
Carothers, Z. B., & Duckett, J. G. 1980. The bryophyte spermatozoid: a source of new phylogenetic
information. Bull. Torrey Bot. Club 107: 281-297.
Carothers, Z. B., & Kreitner, L. 1968. Studies of spermatogenesis in the Hepaticae. II. Blepharoplast

M. Shimamura-26
BSJ- Revi ew 3: 109 ( 2012)

植物科学最前線 3: 110 ( 2012)

Structure in the spermatid of Marchantia. J. Cell Biol. 36: 603-616.
Choi, S. S., Bakalin V. A., & Sun, B. Y. 2010. Unrecorded liverwort species from Mt. Deogyu, Korea. Kor.
J. Plant Tax. 40: 218-225.
Crandall-Stotler, B. 1981. Morphology/anatomy of hepatics and anthocerotes. In: Schultze-Motel W. (ed.)
Advances in Bryology I. pp. 315-398. J. Cramer, Vaduz.
Crandall-Stotler, B., & Stotler. R. E. & Long, D. G. 2009. Phylogeny and Classification of the
Marchantiophyta. Edinb. J. Bot. 66: 155-198.
Dillenius, J. J. 1741-1742. Historia muscorum. Theatro Sheldoniano, Oxford, UK.
Duckett, J. G., & Pressel, S. 2009. Extraordinary features of the reproductive biology of Marchantia atThursley NNR. Field Bryol. 97: 2-11.
Duckett, J. G.. & Renzaglia, K. S. 1993. The reproductive biology of the liverwort Blasia pusilla L. J. Bryol.
17: 541-552.
Equihua, C. 1987. Splash-cup dispersal of gemmae in the liverwort Marchantia polymorpha. Cryptogamie
Bryologie-Lichénologie 8: 199-217.
Forrest, L. L., Davis, E. C., Long, D. G., Crandall-Stotler, B. J., Clark, A., & Hollingsworth. M. L. 2006.
Unraveling the evolutionary history of the liverworts (Marchantiophyta) – Multiple taxa, genomes, andanalyses. Bryologist 109: 303-334.
Graham, L. E., Wilcox, L. W., Cook, M. E., & Gensel, P. G. 2004. Resistant tissues of modern
marchantioid liverworts resemble enigmatic Early Paleozoic microfossils. Proc. Natl. Acad. Sci. USA.
101: 11025-11029.
He-Nygrén, X., Juslén, A., Ahonen, I., Glenny, D., & Piippo, S. 2006. Illuminating the evolutionary historyof liverworts (Marchantiophyta)—towards a natural classification. Cladistics 22: 1-31.
Hofmeister, W. 1851. Vergleichende Untersuchungen der Keimung, Entfaltung und Fruchtbildung hbherer
Kryptogamen und der Samenbildung der Gymnospermen. Leipzig.
Inoue, H. 1960. Studies in spore germination and the earlier stages of gametophyte development in the
Marchantiales. J. Hattori Bot. Lab. 23:148-191.
Izumi, Y., & Ono, K. 1999. Changes in plastid DNA content during the life cycle of the hornwort Anthoceros
punctatus L. Cytologia 64: 37-44
Jankowiak-Siuda, K., Pacak, A., Odrzykoski, I., Wyatt, R. & Szweykowska-Kulińska, Z. 2008. Organellar
inheritance in the allopolyploid mosses species Plagiomnium medium and Plagiomnium curvatulum.
Taxon 57: 145-152.
ȷǰɠ©ĒǗ̕̚ 2012. ǅǚɄt^͌xkh͌v`wfp|. ˌʼ͌ɣɲ 10: 193–210.
üŬŠƼ 1987. r~kh?Ľɮǅǚ=Ʉ-.ǅǚˌʼ͌ō¼¼ĳ 4: 116-117.
üŬŠƼ 1977. r~kh?ʯ͕ȷ:¹ǯ͛ǅǚˌʼ͌ō¼¼ĳ 2: 34-36.
Kammerling, Z. 1897. Zur Biologic und Physiologie der Marchantiaceen. Flora 84: 1-68.
Kny, L. 1890. Bau und Entwicklung von Marchantia polymorpha L. Botanische Wandtafeln 8: 364-401.
Paul Parey, Berlin.
ȏâŌ©ɢũÞſĒ#8ǁ+{Ǳȹ:+8?ʼ͌r~kh?ȺƑBSJ Review 3:

M. Shimamura-27
BSJ- Revi ew 3: 110 ( 2012)

植物科学最前線 3: 111 ( 2012)

58-70.
Konstantinova, N. A., & Bakali, V. A. 2009. Checklist of liverworts (Marchantiophyta) of Russia. Arctoa
18: 1-64.
Kutschera, U., Thomas, J., & Hornschuh, M. 2007. Cluster formation in liverwort- associatedmethylobacteria and its implications. Naturwissenschaften 94: 687-692.
Leitgeb, H., 1880. Die Athemöffnung der Marchantiaceen. Sitz-Ber. d. Kaiserl. Akademie d. Wiss. 81:
40-54.
Linnaeus, C. 1753 Species plantarum. Ed. 1. Salvius, Stockholm.
Ligrone, R., Carafa, A., Lumini, E., Bianciotto, V., Bonfante, P., & Duckett, J. G. 2007. Glomeromycotean
associations in liverworts: a molecular, cellular, and taxonomic analysis. Amer. J. Bot. 94: 1756-1777.
Long, D. G. 2006. New higher taxa of complex thalloid liverworts (Marchantiophyta –Marchantiopsida).
Edinb. J. Bot. 63: 257-262.
McConaha, M. 1941. Ventral structures effecting capillarity in the Marchantiales. Amer. J. Bot. 28:
301-306.
McDaniel, S. F, Willis, J. H., & Shaw, A. J. 2007. A linkage map reveals a complex basis for segregation
distortion in an interpopulation cross in the moss Ceratodon purpureus. Genetics 176: 2489-2500.
Meyer, M., Seibt, U. & Griffiths, H. 2008. To concentrate or ventilate? Carbon acquisition, isotope
discrimination and physiological ecology of early land plant life forms. Phil. Trans. R. Soc. B. 363:
2767-2778
Moser, J. W., & Kreitner, G. L. 1970. Centrosome structure in Anthoceros laevis and Marchantia
polymorpha. J Cell Biol. 44: 454-458.
Nakazato, T., Kadota, A., & Wada, M. 1999. Photoinduction of spore germination in Marchantia
polymorpha L. is mediated by photosynthesis. Plant Cell Physiol. 40: 1014-1020.
O'Hanlon, M. E. 1926. Germination of the spores and early stages in the development of the gametophyte ofMarchantia polymorpha. Bot. Gaz. 82: 215-222.
Pacak, A. & Szweykowska-Kulińska, Z. 2003. Organellar inheritance in liverworts: an example of Pellia
borealis. J Mol Evol. 56:11-7.
Parihar, N. S. 1956. An Introduction to Embryophyta. Volume I. Bryophyta, Central Book Depot.
Allahabad, India.
Paton, J. A. 1999. The Liverwort Flora of the British Isles. Harley Books, Colchester. UK.
Perold, S. M. 1999. Flora of Southern Africa. Hepatophyta. Part 1: Marchantiopsida. In: O.A.Leistner (ed.),Flora of southern Africa:1-252. Botanical Research Institute, Pretoria.
Qiu, Y.-L., Li, L., Wang, B., Chen, Z.. Knoop, V.. Groth-Malonek, M., Dombrovska, O., Lee, J., Kent. L.,
Rest, J., Estabrook, G. F., Hendry, T. A., Taylor, D. W., Testa, C. M., Ambros, M., Crandall-Stotler, B.,
Duff, R. J., Stech, M., Frey, W., Quandt, D., & Davis C. C. 2006. The deepest divergences in land plants
inferred from phylogenomic evidence. Proc. Natl. Acad. Sci. USA 103: 15511-15516.
Qiu, Y.-L., Li, L., Wang, B., Chen, Z., Dombrovska, O., Lee, J., Kent, L., Li, R., Jobson, R. W., Hendry, T.
A., Taylor, D. W., Testa, C. M., & Ambros, M. 2007. A nonflowering land plant phylogeny inferred

M. Shimamura-28
BSJ- Revi ew 3: 111 ( 2012)

植物科学最前線 3: 112 ( 2012)

from nucleotide sequences of seven chloroplast, mitochondrial, and nuclear genes. Int. J. Plant Sci. 168:691-708.
Renzaglia, K.S., & Garbary, D. J. 2001. Motile gametes of land plants: diversity, development, andevolution. Crit Rev Plant Sci. 20:107-213
Schiffner, V. 1909 Studien über die Rhizoiden der Marchantiales. Ann. Jard. Bot. Buitenz. 2 ser., Suppl. 3:
473-492.
Schuster, R. 1983. Phytogeography of the Bryophyta. In New manual of bryology. Edited by R.M. Schuster.
Hattori Bot. Lab., Nichinan, Japan.
Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America, Vol. 6. Field Museum of Natural
History. Chicago.
Shimamura, M., Fukushima, H., Yamaguchi, T., & Deguchi, H. 1999. Behavior of plastid and plastid DNA
during spermiogenesis in Dumortiera hirusta. Bryol. Res. 7: 201-204.
Shimamura, M., Brown, R. C., Lemmon, B. E., Akashi, T., Mizuno, K., Nishihara, N. Tomizawa K. I.,
Yoshimoto, K., Deguchi, H., Hosoya , H., Horio, T., & Mineyuki, Y. 2004. γ-Tubulin in basal land
plants: characterization, localization, and implication in the evolution of acentriolar microtubuleorganizing centers. Plant Cell 16: 45-59.
Shimamura, M., Furuki, T., & Deguchi, H. 2005. Sporophyte anatomy of Cavicularia densa Steph.
(Blasiaceae). Bryologist. 108: 420-426.
ūǜȀǻĒǗ̕̚çđăñ 2006. _pr~kh?ʪňÄ=%UƆƚōɒɣɲ ͛Hikobia 14:
399-402.
Shimamura, M., Yamaguchi, T., & Deguchi, H. 2008. Airborne sperm of Conocephalum conicum(Conocephalaceae). J. Plant Res. 121: 69-71.
Shimamura, M., Itouga, M., & Tsubota, H. 2012. Evolution of apolar sporocytes in marchantialeanliverworts: implications from molecular phylogeny. J. Plant Res. 125: 197-206.
Stotler, R. 1994. [Review of the book Marchantia L. The European and African Taxa. Bryophytrum
Bibliotheca 45 by Bishler, H.] Bryologist 97: 351-352.
Suire, C., Bouvier, F., Backhaus, R. A., Bégu, D., Bonneu, M., & Camara, B. 2000. Cellular localization of
isoprenoid biosynthetic enzymes in Marchantia polymorpha. uncovering a new role of oil bodies. Plant
Physiol. 124: 971-978.
Völker, H, 1972. Licht- und elektronenmikroskopische Untersuchungen zur Cytokinese an jungenBrutkörpern vonMarchantia polymorpha L. Protoplasma, 75: 139-153.
Walker, R., & Pennington, W. 1939. The movements of the air pores of Preissia quadrata (Scop.). New
Phytologist 38: 62-68.
Walton, J. 1925. Carboniferous bryophytes. I. Hepaticae. Ann. Bot. 39: 563-72.
Wellman, C. H., Osterff, P. L., & Mohiuddin, U. 2003. Fragments of the earliest land plants, Nature 425:282-285.
Wheeler, J. A. 2000. Molecular phylogenetic reconstructions of the marchantioid liverwort radiation.
Bryologist 103: 314-333.

M. Shimamura-29
BSJ- Revi ew 3: 112 ( 2012)

植物科学最前線 3: 113 ( 2012)

Yuasa, A. 1952. Studies in the cytology of Pteridophyta XXX. The behavior of the plastid during theprocess of fertilization. Sci. Pap. Coll. Gen. Edu. Univ. Tokyo 2: 73-79.

M. Shimamura-30
BSJ- Revi ew 3: 113 ( 2012)